Cryptic diversity in the subgenus Oxyphortica (Diptera, Drosophilidae, Stegana)

Sampling and morphological evaluations

Specimens were collected by net sweeping from tussocks or tree trunks along streams in forests. Following collection, the specimens were immediately preserved in 75% ethanol. In the laboratory, all samples collected from the field were initially identified based on morphological characteristics, and then targeted specimens were selected for thorough morphologilical examination. Detailed information is listed in Table 1. Genitalia of the specimens were removed for further identification and photographed using a microscope Mshot Camera (Mshot, China). A small piece of tissue was excised for DNA extraction using a TIANGEN™ DNA Extraction Kit (Tiangen Biotech, Beijing, China). Finally, using a combination of morphological characteristics and molecular data, new species were recognised. Type specimens were air-dried and deposited in the Kunming Institute of Zoology, Chinese Academy of Sciences, Kunming, China (KIZ); the Department of Zoology, Tibet Museum of Natural Science, Lasa, China (TMNS); and the Department of Entomology, South China Agricultural University, Guangzhou, China (SCAU). To unify Drosophilidae terminology and facilitate exchanges between research fields, the male terminalia terminology followed that of Rice et al. (2019).

The electronic version of this article in portable document format will represent a published work according to the International Commission on Zoological Nomenclature (ICZN), and hence the new names contained in the electronic version are effectively published under that Code from the electronic edition alone. This published work and the nomenclatural acts it contains have been registered in ZooBank, the online registration system for the ICZN. The ZooBank LSIDs (Life Science Identifiers) can be resolved and the associated information viewed through any standard web browser by appending the LSID to the prefix http://zoobank.org/. The LSID for this publication is: urn:lsid:zoobank.org:pub:1D19FB6F-DBF2-4982-9C9A-D8FBFC1F3F1E. The online version of this work is archived and available from the following digital repositories: PeerJ, PubMed Central and CLOCKSS.

PCR amplification and sequencing

Two mitochondrial (cytochrome c oxidase subunit I (COI) and NADH dehydrogenase subunit 2 (ND2)) markers were employed for PCR amplification, which involved the following steps: initial pre-denaturation for 3 min at 94 °C, followed by 35 cycles of 30 s at 94 °C, 1 min at 49–56 °C and 1 min at 72 °C, and subsequent post-extension for 5 min at 72 °C. The amplification and sequencing primers are listed in Table 2. Sequencing of the PCR products was performed using an ABI 3700 sequencer; a standard sequencing reaction using a BigDye™ Terminator Kit (Perkin-Elmer, Waltham, MA, USA) was performed with 25 cycles of denaturation for 10 s at 96 °C, annealing for 5 s at 50 °C and extension for 4 min at 60 °C, subsequently followed by purification with 75% isopropyl alcohol. The sequence accession numbers are listed in Table 1.

Phylogenetic analyses

The mitochondrial sequences were aligned separately for each gene using the programme MAFFT v. 7 (http://mafft.cbrc.jp/alignment/server/) and then concatenated (COI + ND2). A total of six data blocks of the concatenated data (i.e., first-, second- and third-codon positions of two mitochondrial genes) were subjected to an evaluation of the best partitioning scheme and substitution models (GTR+I+G: COI^1st + COI^2nd + ND2^1st + ND2^2nd; TIM+I+G: COI^3rd + ND2^3rd) for phylogenetic analysis in PartitionFinder 2 (Lanfear et al., 2017) using the ′greedy′ algorithm and the Bayesian information criterion. Phylogenetic analyses were then performed using Bayesian inference and maximum likelihood (ML) methods. Twelve outgroup species (Leucophenga angusta Okada, 1956; L. quadripunctata (de Meijere, 1908); Parastegana brevivena Chen & Zhang, 2007; Pa. punctalata (Chen & Watabe, 2007); Pseudostegana meiduo Zhang & Chen, 2018; Ps. meiji Zhang & Chen, 2018; S. (Steganina) acantha Wu, Gao & Chen, 2010; S. (St.) ancistrophylla Wu, Gao & Chen, 2010; S. (sensu stricto) apiciprocera Cao & Chen, 2010; S. (s.s.) quadrata Cao & Chen, 2010; S. (Orthostegana) flavicauda Zhang & Chen, 2012 and S. (Or.) hirsutina Zhang & Chen, 2012) were selected from three Steganinae genera (Leucophenga, Parastegana and Pseudostegana) and three closely related subgenera (Stegana, Steganina and Orthostegana), respectively. Several sequences from the Steganinae genera and subgenera as well as from Stegana genus sensu stricto were downloaded from GenBank.

Bayesian inference was performed using MrBayes v.3.2.1 (Huelsenbeck & Ronquist, 2001; Ronquist & Huelsenbeck, 2003) on the CIPRES Science Gateway (http://www.phylo.org; Miller, Pfeiffer & Schwartz, 2010). Two independent runs were implemented in parallel, each with 20,000,000 generations. A sampling frequency of every 1,000 generations was employed, and 5,000 early-phase samples were discarded as burn-in. When the average standard deviation of split frequencies was <0.01, the analysis was considered convergent. The ML analysis was performed using IQ-Tree (Nguyen et al., 2015) under the aforementioned substitution models. Node support was assessed via ultrafast bootstrap (UFBP) analysis with 1,000 replicates. MEGA 5 was used to estimate pairwise genetic distances (p-distance) (Tamura et al., 2011).

Species delimitation

Species delineation was explored using Automatic Barcode Gap Discovery (ABGD; Puillandre et al., 2012) for each of the two single-locus data, and using Bayesian Phylogenetics and Phylogeography (BP&P; Rannala & Yang, 2003; Yang & Rannala, 2010) for the concatenated dataset. The ABGD analysis was performed at the web interface (http://www.abi.snv.jussieu.fr/public/abgd/) with parameters that included P from 0.001 to 0.1, relative gap width (X = 1.0) and the Kimura-2-parameter (K2P; Kimura, 1980) model. The BP&P analysis was implemented in the BP&P v.3.2 programme with the method A11 (Yang, 2015). In the Bayesian analysis, the ancestral population size (θ), root age (τ0) and key priors of the Bayesian species delimitation may strongly affect the posterior probabilities of the speciation modes (Yang & Rannala, 2010); therefore, four different gamma prior (G) combinations were considered for the BP&P coalescent analysis: (1) large ancestral population sizes and deep divergences, θ~G (1, 10) and τ0~G (1, 10); (2) small ancestral population sizes and shallow divergences, θ~G (2, 2,000) and τ0~G (2, 2,000); (3) large ancestral population sizes and shallow divergences, θ~G (1, 10) and τ0~G (2, 2,000); (4) small ancestral population sizes and deep divergences, θ~G (2, 2,000) and τ0~G (1, 10). All analyses were performed twice to confirm consistency. The starting guide tree was first generated using BEAST v.1.10.4 (Suchard et al., 2018) with the GTR + G + I substitution model and the Yule speciation model. We set the length of the chain to 20,000,000, sampled every 2,000 generations and discarded the first 25% as burn-in. The convergence was evaluated using Tracer v.1.7 (Rambaut et al., 2018). When the effective sample size was >200, the analysis was considered convergent.

Taxonomy

Subgenus Stegana (Oxyphortica) Duda, 1923

Phortica (Oxyphortica) Duda, 1923: 34. Type species: Drosophila convergens de Meijere, 1911.

Chaetocnema Duda, 1926: 243. Type species: Chaetocnema poeciloptera Duda, 1926 (= Orthostega nigripennis Hendel, 1914).

Stegana (Oxyphortica): Okada, 1971: 90; Okada, 1978: 398; Toda & Peng, 1992: 210; Sidorenko, 1998: 297; Xu et al., 2007: 43; Wang et al., 2010: 566.

Diagnosis. Mesoscutum with ca. 12 irregular rows of acrostichal setulae; first tarsomere of foreleg at least basally and apically, each with two or three black, long, spine-shaped setae on ventral surface; pregonites (parameres in McAlpine, 1981) mostly inconspicuous or absent, if present, usually symmetrical; aedeagus diverse, not annular, occasionally absent; phallapodeme (aedeagal apodeme in McAlpine, 1981) without process subapically.

Description. Eyes brownish red. Ocellar triangle black. Postocellar setae divergent, absent in the convergens group. Frons brown to dark brown, slightly narrowed below, mostly with sparse interfrontal setulae. Pedicel yellow, with one long seta and several minute setulae; first flagellomere mostly greyish yellow. Face brown. Clypeus narrow, yellow to black. Gena dark brown. Palpus mostly yellow and slender, occasionally dark brown, with several setulae. Mesoscutum yellow to brown, with variable pattern. Postpronotal lobe yellow, with one long seta. Pleuron with a brown longitudinal stripe above (running from propleuron to base of haltere). Katepisternum yellow to brown, with two long and some small setae. Wing greyish brown to dark brown; costal vein between R₂₊₃ and R₄₊₅ with five to eight peg-like spinules on ventral surface; vein M₁ distally variably convergent to vein R₄₊₅. Haltere brownish on stem, greyish yellow on knob. Legs mainly yellow, with occasional brown tinge or darker brown markings; forefemur with a row of long setae on ventral surface; midtibia with two strong setae on dorsal surface (as Fig. 3D in Zhang et al., 2012). Abdominal tergites yellow to black; sternites mostly yellow. Male terminalia: Cercus mostly lacking pubescence. Surstylus with or without diverse prensisetae. Hypandrial phragma (Hypandrium in McAlpine, 1981) mostly expanded anteriorly. Aedeagal sheaths (Gonopods in McAlpine, 1981) fused, mostly basolaterally extended, fused with lateral gonocoxite (hypandrium in McAlpine, 1981).

Seventeen new species that resemble seven known species (S. (O.) crassiforcipata Wang, Gao & Chen, 2010; S. (O.) curvata Wang, Gao & Chen, 2010; S. (O.) enigma Sidorenko, 1998; S. (O.) monoacantha Wang & Chen, 2010; S. (O.) subconvergens Okada, 1988; S. (O.) wanglei Wang, Gao & Chen, 2010 and S. (O.) wuliangi Wang, Gao & Chen, 2010) in some morphological characteristics (Figs 3–5) (e.g., foretarsi mostly with fringe-like setae on anterior surface; surstylus narrowed and curved, dorsally mostly with one to two stout (occasionally a row of small) prensisetae (Figs 6–22B)) are described.

Stegana (Oxyphortica) amphigya Wang & Chen, sp. nov.

urn:lsid:zoobank.org:act:E6BE14A6-5E2B-4DAD-87D6-2819DFAE5202

(Figs 3D, 5D, 6)

Type specimens. Holotype, ♂ (SCAU, no. 111283), CHINA: Mengdong, Cangyuan, Yunnan, 23°100′8″N, 99°13′52″E, 1,320–1,400 m a.s.l. (above sea level), 6.v.2016, ex tussock (J Huang). Paratypes, CHINA: 5♂, 1♀ (SCAU, nos 111284–89), ex tussocks and tree trunks, same data as holotype (Y. Liu, Y. Wang, L Zhu); 1♂, 1♀ (KIZ, nos 0090651, 52), Baihualing, Baoshan, Yunnan, 25°46′02″N, 98°43′44″E, 1,400–1,600 m a.s.l., 23.viii.2013, ex tussock (J. Gao); 6♂, 8♀ (SCAU, nos 111290–303), Husa, Longchuan, Yunnan, 24°27′37″N, 97°45′9″E, 1,230–1,340 m a.s.l., 20.viii.2016, ex tree trunks (H. Chen, L Gong, Y. Liu).

Diagnosis. Foretarsi second to fourth subapically, each with one small, suberect, fringe-like seta on anterior surface (Fig. 3D). Surstylus acute ventrally, with six prensisetae on dorstal half (Fig. 6B); aedeagal sheaths submedially with two pairs of prensisetae and four pairs of setae, distolaterally strongly, elongated to semiring-shaped, with a few setae distally and one small prensisetae on one side (Figs. 6C, 6D).

Description. Arista with four to six dorsal and three to five ventral branches. Clypeus yellowish brown. Mesoscutum and scutellum yellow to yellowish brown. Katepisternum brown above, yellow below. Abdominal first tergite yellow; second to sixth tergites mostly dark brown, yellow medially, narrowly yellow bands on anterior margins on fourth and fifth tergites (Fig. 5D). Sternites mostly brown, yellow on second. Male terminalia (Fig. 6): Epandrium with ca. 20 setae on each posterior portion. Surstylus with a row of ca. 11 setae. Pregonites small and inconspicuous. Aedeagus with pubescence distally.

Measurements. Body length (BL) = 2.83 mm in holotype (5♂ and 5♀ paratypes: 2.16–2.67 in ♂, 2.53–3.20 in ♀), thorax length (THL) = 1.17 mm (0.93–1.17, 1.07–1.37), wing length (WL) = 2.33 mm (1.83–2.23, 2.17–2.70), wing width (WW) = 1.07 mm (0.93–1.07, 0.93–1.30).

Etymology. From the Greek word: amphigyos, referring to the surstylus pointed on both sides.

Stegana (Oxyphortica) armillata Wang & Chen, sp. nov.

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(Figs. 3E, 5E, 7)

Type specimens. Holotype, ♂ (SCAU, no. 111276), CHINA: Dashuigou, Lüchun, Yunnan, 22°57′19″N, 101°57′04″E, 1,310 m a.s.l., 2.viii.2016, ex tussock (H. Chen). Paratypes, CHINA: 6♂, 3♀ (2♂, 1♀ in KIZ, nos 0090648–50; 4♂, 2♀ in SCAU, nos 111277–82), Sanmeng, Lüchun, Yunnan, 22°53′25″N, 102°19′00″E, 1,560 m a.s.l., 1.viii.2016, ex tussocks (H. Chen, L Gong, Y. Liu).

Diagnosis. This species is very similar to S. (O.) amphigya sp. nov. in male terminalia, but can be distinguished by the foretarsi second to fourth subapically lacking small, suberect, fringe-like seta on anterior surface (Fig. 3E); surstylus round ventrally, with seven small prensisetae on dorstal 3/4 (Fig. 7B).

Description. Arista with four to six dorsal and three to four ventral branches. Clypeus brown. Mesoscutum and scutellum yellow to brownish yellow. Katepisternum brownish above, yellow below. Abdominal first tergite yellow; second and third tergites yellow medially, brown laterally; fourth to sixth tergites mostly brown, yellow on anterior margins (Fig. 5E). Sternites yellow, with narrowly brown band on lateral margins of fourth to sixth. Male terminalia (Fig. 7): Epandrium with ca. 21 setae on each posterior portion. Surstylus with a row of ca. 12 setae. Aedeagal sheaths submedially with two pairs of prensisetae and five pairs of setae, elongated to semiring-shaped, distally with one small prensisetae and a few setae on each side. Pregonites small, inconspicuous. Aedeagus with pubescence distally.

Measurements. BL = 2.43 mm in holotype (5♂ and 3♀ paratypes: 2.40–3.33 in ♂, 2.60–3.38 in ♀), THL = 1.07 mm (0.93–1.33, 1.07–1.51), WL = 2.17 mm (1.87–2.57, 2.20–2.83), WW = 1.03 mm (0.87–1.23, 0.97–1.37).

Etymology. From the Latin word: armillatus, referring to the semiorbicular surstylus.

Stegana (Oxyphortica) ashima Zhang & Chen, sp. nov.

urn:lsid:zoobank.org:act:5F7EECB0-12D6-419F-90BB-697D9BF2C455

(Figs. 3F, 5F, 8)

Type specimens. Holotype, ♂ (SCAU, no. 121350), CHINA: Cangshan, Yangbi, Yunnan, 25°38′22″N, 100°01′55″E, 1,800 m a.s.l., 26.viii.2011, ex tussock (H. Chen). Paratypes, CHINA: 4♂, 4♀ (2♂, 1♀ in KIZ, nos 0090645–47; 2♂, 3♀ in SCAU, nos 121351–55), same data as holotype (H. Chen, J. Gao).

Diagnosis. Epandrium with ca. six long and nine short setae on posterior portion per side (Fig. 8A); pregonites asymmetric (Fig. 8C); phallapodeme anterodistally developed, annularly fused and deeply concave posteromedially (Fig. 8C, phap); aedeagus with a row of setae on apical margin (Fig. 8C).

Description. Arista with four to six dorsal and three to four ventral branches. Clypeus black. Palpus brown. Mesoscutum brownish yellow to brown. Katepisternum brown, sometimes yellow below. Scutellum brown. Foretarsi first to third with ca. 12 valgus fringe-like setae on anterior surface (Fig. 3F). Abdominal first to third tergites yellow medially, brown to black laterally; fourth to sixth tergites mostly brown, yellow dorsally along anterior margins (Fig. 5E). Sternites brownish yellow. Male terminalia (Fig. 8): Surstylus with one stout prensisetae subdorsally, a row of ca. 10 setae medially and five setae ventrally. Hypandrial phragma narrowed anteriorly. Aedeagal sheath slender.

Measurements. BL = 2.30 mm in holotype (4♂ and 3♀ paratypes: 1.83–2.50 in ♂, 2.07–2.17 in ♀), THL = 0.93 mm (0.83–1.07, 0.83–0.90), WL = 2.00 mm (2.00–2.50, 2.00–2.30), WW = 0.83 mm (0.83–1.00, 0.83–0.87).

Etymology. Name of a beautiful woman from legend of the Yi ethnic group in Yunnan, China.

Stegana (Oxyphortica) bawo Zhang & Chen, sp. nov.

urn:lsid:zoobank.org:act:7EF83FD7-92D3-4831-BBA6-3ACDF305E9D0

(Figs. 3G, 5G, 9)

Type specimens. Holotype, ♂ (SCAU, no. 121365), CHINA: Hutiaoxia, Xianggelila, Yunnan, 27°11′24″N, 100°02′54″E, 1,700 m a.s.l., 22.viii.2011, ex tussock (H. Chen). Paratype, CHINA: 1♀ (SCAU, no. 121366), Zhengxing, Jinggu, Yunnan, 23°19′54″N, 100°57′45″E, 1,100–1,300 m a.s.l., 24.vii.2009, ex tussock (L Wang).

Diagnosis. This species is very similar to S. (O.) mouig sp. nov. in the shape of phallapodeme anterodistally (Fig. 15C), but can be distinguished by the surstylus with one stout prensisetae dorsally (Fig. 9B) and aedeagus entirely membranous (Fig. 9C).

Description. Arista with seven to eight dorsal and four ventral branches. Clypeus dark brown. Mesoscutum mostly yellow, sometimes light brown laterally. Katepisternum brown above, yellow below. Scutellum brownish yellow. Foreleg first tarsus with ca. seven fringe-like setae; first to fourth tarsi with ca. 11 setae and subapically each with one valgus fringe-like setae on anterior surface (Fig. 3G). Abdominal first and second tergites mostly yellow, dark brown on second tergite posterior margin; third yellow medially, dark brown laterally; fourth to sixth dark brown to black (Fig. 5G). Sternites yellow on first to third, dark brown to black on fourth to sixth. Male terminalia (Fig. 9): Epandrium with ca. 22 setae on each posterior portion. Surstylus with two irregular rows of numerous setae medially and a few setae ventrally. Pregonites absent. Aedeagus distally plicated. Aedeagal sheath slender.

Measurements. BL = 3.20 mm in holotype (1♀ paratype: 3.00 mm), THL = 1.27 mm (1.27), WL = 2.80 mm (2.73), WW = 1.20 mm (1.27).

Etymology. The name denotes “dauntless” in Tibetan.

Stegana (Oxyphortica) crypta Wang & Chen, sp. nov.

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(Figs. 3H, 5H, 10)

Type specimen. Holotype, ♂ (SCAU, no. 111305), CHINA: Dashuigou, Lüchun, Yunnan, 22°57′19″N, 101°57′04″E, 1,310 m a.s.l., 2.viii.2016, ex tussock (H. Chen).

Diagnosis. This species differs from other species in having the surstylus very narrow, with two small prensisetae dorsally (Fig. 10C); aedeagus with numerous serrated processes on lateral margins (Figs. 10C, 10D).

Description. Arista with six dorsal and four ventral branches. Clypeus brown. Mesoscutum yellow, slightly brownish laterally and posteriorly. Scutellum brownish yellow. Katepisternum entirely yellow. Foretarsi second to fourth each with one small fringe-like seta on anterior surface (Fig. 3H). Abdominal tergites all yellow medially, brown laterally (Fig. 5H); sternites yellow. Male terminalia (Fig. 10): Epandrium with ca. 22 setae on posterior portion per side. Surstylus with two irregular rows of setae medially, and ca. six setae ventrally. Pregonites present. Aedeagal sheaths vertical, elongated distolaterally. Aedeagus with a row of setae apically.

Measurements. BL = 2.67 mm, THL = 1.10 mm, WL = 2.10 mm, WW = 1.00 mm.

Etymology. From the Greek word: cryptos, meaning that the fly hides in the mountains.

Stegana (Oxyphortica) gelea Zhang & Chen, sp. nov.

urn:lsid:zoobank.org:act:91334019-DC7F-4922-9D95-25B8C7CD64C2

(Figs. 3I, 5I, 11)

Type specimens. Holotype, ♂ (SCAU, no. 121356), CHINA: Tongmai, Bomi, Xizang, 30°06′10″N, 95°04′48″E, 2,080 m a.s.l., 11.x.2010, ex tussock (J. Gao). Paratypes, CHINA: 3♂, 3♀ (1♂, 1♀ in KIZ, nos 0090653–54; 2♂, 2♀ in SCAU, nos 121357–60), 9.x.2010, same data as holotype (J. Gao); 2♂, 2♀ (TMNS), Beibeng, Motuo, Xizang, 29°14′36″N, 95°19′56″E, 1,100 m a.s.l., 27.ix.2010, ex tussocks (Y. Su, L Wang, L Wu).

Diagnosis. This species is very similar to S. (O.) wanhei sp. nov. in the male terminalia structure, but can be distinguished from the latter by the apical processes of aedeagus and aedeagal sheaths are nearly equilong (Figs. 11C, 11D), foreleg first tarsus subapically to apically with three recurvate fringe-like setae on anterior surface (Fig. 3I).

Description. Arista with six dorsal and four to five ventral branches. Clypeus black. Mesoscutum brownish yellow. Katepisternum brown anteriorly, yellow posteriorly. Scutellum brown, yellow along margin. Abdominal first tergite yellow; second to fourth tergites dark brown, yellow anteromedially; fifth and sixth tergites black, yellow on anterior margins (Fig. 5I). Sternites entirely yellow. Male terminalia (Fig. 11): Epandrium with ca. 30 setae on posterior portion per side. Surstylus with two pointed prensisetae dorsally and two irregular rows of numerous setae medially, elongated and curved ventrally. Pregonites small. Aedeagus with several setae medially, mediodorsally fused with aedeagal sheaths, bifurcated distally. Aedeagal sheaths bifurcated distally.

Measurements. BL = 3.34 mm in the holotype (4♂ and 5♀ paratypes: 3.16–3.56 in ♂, 3.00–4.00 in ♀), THL = 1.64 mm (1.48–1.74, 1.48–1.88), WL = 3.56 mm (2.88–3.44, 2.88–3.80), WW = 1.76 mm (1.32–1.60, 1.28–2.90).

Etymology. The name means “lucky place” in Tibetan.

Stegana (Oxyphortica) hengduanmontana Zhang & Chen, sp. nov.

urn:lsid:zoobank.org:act:C2BA7D62-428B-4FA7-9507-24AC574813E8

(Figs. 3J, 5J, 12)

Type specimens. Holotype, ♂ (SCAU, no. 122227), CHINA: Yixiang, Pu′er, Yunnan, 22°44′19″N, 101°07′01″E, 1,420 m a.s.l., 12.xii.2012, ex tussock (J. Gao). Paratypes, CHINA: 1♀ (SCAU, no. 122228), same data as holotype; 3♂ (SCAU, nos 111273–75), 28.iii.2013, same data as holotype; 1♂, 1♀ (SCAU, nos 111719, 20), Mengma, Menglian, Yunnan, 22°12′59″N, 99°22′51″E, 1,060 m a.s.l., 14.iv.2018, ex tussock (L Gong).

Diagnosis. This species differs from other species in having the foreleg first tarsus with ca. 12 small, fringe-like setae on anterior surface (Fig. 3J); aedeagus and aedeagal sheaths strongly sclerotized and glabrous (Figs. 12C, 12D).

Description. Arista with six dorsal and four ventral branches. Clypeus brown. Mesoscutum and scutellum brownish yellow. Katepisternum brown anteriorly, yellow posteriorly. Abdominal first tergite yellow; second and third tergites yellow medially, brown laterally; fourth to sixth mostly brown, with yellow anteromedially (Fig. 5J). Sternites yellow. Male terminalia (Fig. 12): Epandrium with two irregular rows of ca. 20 setae along each posterior margin. Cercus with pubescence. Surstylus with two stout prensisetae dorsally, pubescence medially, and ca. five setae ventrally. Pregonites present. Aedeagus apically bifurcated and curved. Aedeagal sheaths anterolaterally elongated, curved and fused with phallapodeme, posteriorly protruded.

Measurements. BL = 2.80 mm in holotype (1♀ paratype: 2.60 mm), THL = 1.13 mm (1.13), WL = 2.53 mm (2.47), WW = 1.00 mm (1.07).

Etymology. A combination of the type locality + the Latin word montanus (= montane), referring to the specimens were collected in Hengduan Mountains.

Stegana (Oxyphortica) jinmingi Zhang & Chen, sp. nov.

urn:lsid:zoobank.org:act:D744D8E0-9475-474F-971F-8EDE39E9C756

(Figs. 4A, 5K, 13)

Type specimens. Holotype, ♂ (SCAU, no. 121251), CHINA: Caiyanghe Forest Park, Pu′er, Yunnan, 27°11′12″N, 100°02′59″E, 900–1,400 m a.s.l., 27.vii.2009, ex tussock (L Wu). Paratypes, CHINA: 1♂, 1♀ (SCAU, nos 121252, 53), same data as holotype; 2♀ (KIZ, nos 0090596, 97), Yixiang, Pu′er, Yunnan, 22°44′19″N, 101°07′01″E, 1,420 m a.s.l., 28.iii.2013, ex tussock (J. Gao); 2♂, 5♀ (SCAU, nos 121254, 55, 121495, 122706–09), Hesong, Menghai, Yunnan, 21°50′08″N, 100°05′51″E, 1,700–1,900 m a.s.l., 27.iii.2011, 12.v.2012, ex tussocks (J. Gao, J. Lu, Y. Su, L Wang, L Wu, S. Yan); 4♂ (SCAU, nos 122710–13), Husa, Longchuan, Yunnan, 24°27′37″N, 97°45′9″E, 1,230–1,340 m a.s.l., 20.viii.2016, ex tussocks (H. Chen, L Gong, Y. Liu); 1♂, 6♀ (SCAU, nos 111259–65), Qimaba, Lüchun, Yunnan, 22°49′03″N, 102°17″39″E, 1,280 m a.s.l., 30.x.2016, ex tussocks (H. Chen); 6♂ (SCAU, nos 111714, 15, 111017–20), CHINA: Botanic Garden, Ruili, Yunnan, 24°01′12″N, 97°51′33″E, 1,170 m a.s.l., 6.xi.2017, ex tussocks (H. Chen, L Gong, B. Li).

Diagnosis. This species is very similar to S. (O.) wanhei sp. nov. in the male terminalia, but can be distinguished from the latter by the apical processes of aedeagal sheaths slightly curved in lateral view (Fig. 13D); foretarsi first and second with two rows of ca. 10 long, recurvate fringe-like setae; first to fourth subapically each with one valgus fringe-like seta on anterior surface (Fig. 4A).

Description. Arista with six dorsal and four ventral branches. Clypeus brown. Mesoscutum yellow, sometimes light brown laterally. Katepisternum yellow, brown on anterior corner. Scutellum brownish yellow. Abdominal first tergite yellow; second and third tergites yellow medially, dark brown laterally; fourth and fifth mostly dark brown, yellow anteromedially; sixth dark brown (Fig. 5K). Sternites yellowish brown. Male terminalia (Fig. 13): Epandrium with ca. 27 setae on posterior portion per side. Surstylus with two pointed prensisetae dorsally and numerous setae ventrally, elongated and slightly curved ventrally. Pregonites absent. Aedeagus with several setae medially, mediodorsally fused with aedeagal sheaths, bifurcated distally. Aedeagal sheaths bifurcated distally.

Measurements. BL = 2.50 mm in holotype (1♂ paratype: 2.80 mm), THL = 1.34 mm (1.37), WL = 2.18 mm (2.33), WW = 1. 00 mm (1.13).

Etymology. Patronym after the collector Mr. Jinming Lu (SCAU).

Stegana (Oxyphortica) mengbalanaxi Zhang & Chen, sp. nov.

urn:lsid:zoobank.org:act:35238DA6-11C8-49AC-8BA2-B4511D549F65

(Figs. 4B, 5L, 14)

Type specimens. Holotype, ♂ (SCAU, no. 121367), CHINA: Baihualing, Baoshan, Yunnan, 25°46′02″N, 98°43′44″E, 1,400–1,600 m a.s.l., 14.vi.2011, ex tussock (J. Gao). Paratypes, CHINA: 1♀ (SCAU, no. 121368), same data as holotype (H. Chen); 1♂, 1♀ (SCAU, nos 124776, 77), 21.vi.2013, same data as holotype (Y. Wang); 1♂ (SCAU, no. 124175), Husa, Longchuan, Yunnan, 24°27′37″N, 97°45′9″E, 1,230–1,340 m a.s.l., 20.viii.2016, ex tree trunk (H. Chen).

Diagnosis. This species somewhat resembles S. (O.) yangjin sp. nov. in the male terminalia (Figs. 14, 21), but can be distinguished from the latter by the foretarsi first and second with two to three rows of ca. 22 long, recurvate fringe-like setae; first to fourth apically each with one short, valgus fringe-like seta on anterior surface (Fig. 4B).

Description. Arista with seven to eight dorsal and four to five ventral branches. Clypeus black. Palpus brown. Mesoscutum brownish to brown. Katepisternum dark brown above, yellow below. Scutellum dark brown. Abdominal first tergite yellow; second to sixth tergites mostly dark brown, with yellow patches submedially (Fig. 5L). Sternites brown. Male terminalia (Fig. 14): Epandrium with ca. 24 setae on posterior portion per side. Surstylus with two prensisetae dorsally, pubescence and setae submedially and ventrally. Pregonites absent. Aedeagus with several setae medially and numerous minute serrated processes laterally, fused with aedeagal sheaths dorsomedially. Aedeagal sheaths curved posteriorly, acute apically.

Measurements. BL = 3.00 mm in holotype (1♂ paratype: 2.33 mm), THL = 1.20 mm (1.00), WL = 2.67 mm (2.00), WW = 1.40 mm (0.93).

Etymology. The name means “dreamlike homeland”, from the language of the Dai ethnic group in Xishuangbanna, Yunnan, China.

Stegana (Oxyphortica) mouig Zhang & Chen, sp. nov.

urn:lsid:zoobank.org:act:7991CFC0-2FCC-426C-AEAE-5CA088F2EB46

(Figs. 4C, 5M, 15)

Type specimens. Holotype, ♂ (SCAU, no. 121369), CHINA: Yixiang, Pu′er, Yunnan, 22°44′19″N, 101°07′01″E, 1,420 m a.s.l., 2.x.2011, ex tussock (H. Chen). Paratypes, CHIAN: 1♂, 1♀ (SCAU, nos 111134, 121370), same data as holotype; 11♂, 4♀ (5♂, 2♀ in KIZ, nos 0090598–604; 6♂, 2♀ in SCAU, nos 111135–42), 28–30.iii.2013, 28.v.2015, ex tussocks and tree trunks, same data as holotype (H. Chen, J. Gao); 3♂ (SCAU, nos 122931, 123019, 20), Wuliangshan, Jingdong, Yunnan, 24°17′27″N, 100°19′48″E, 1,700–2,100 m a.s.l., 1.ix.2014, ex tussock (J. Gao); 35♂, 25♀ (10♂, 10♀ in KIZ, nos 0090605–24; 15♂, 15♀ in SCAU, nos 111143–69, 111192–201, 111721–23), Mengdong, Cangyuan, Yunnan, 23°100′8″N, 99°13′52″E, 1,320–1,400 m a.s.l., 6.v.2016, 19.iv.2019, ex tussocks and tree trunks (H. Chen, L Gong, J Huang, Y Lin, Y. Liu, Y. Wang, L Zhu); 14♂, 2♀ (SCAU, nos 111170–85), Dashuigou, Lüchun, Yunnan, 22°57′19″N, 101°57′04″E, 1,310 m a.s.l., 2.viii.2016, ex tussocks and tree trunks (H. Chen, L Gong, Y. Liu); 5♂ (SCAU, nos 111186–91), Husa, Longchuan, Yunnan, 24°27′37″N, 97°45′9″E, 1,230–1,340 m a.s.l., 20.viii.2016, ex tussocks (H. Chen, L Gong, Y. Liu).

Diagnosis. This species differs from S. (O.) bawo sp. nov. in having the surstylus with two stout prensisetae dorsally (Fig. 15B); aedeagus round distally, slightly sclerotized medially (Fig. 15C).

Description. Arista with four to six dorsal and three to four ventral branches. Clypeus brown. Mesoscutum yellow, with brown longitudinal stripes. Katepisternum yellow. Scutellum brown, yellow along margin. Foretarsi first to fourth with a row of ca. 13 long, recurvate fringe-like setae, and subapically each with one short, valgus fringe-like seta on anterior surface (Fig. 4C). Abdominal first tergite yellow; second tergite yellow medially, dark brown laterally; third tergite mostly dark brown, with one pair of yellow patches medially; fourth to sixth dark brown (Fig. 5M). Sternites yellow. Male terminalia (Fig. 15): Epandrium with ca. 19 setae on posterior portion per side. Surstylus with numerous setae submedially and ventrally. Aedeagal sheath slender.

Measurements. BL = 2.67 mm in holotype (5♂ and 5♀ paratypes: 2.33–3.00 in ♂, 2.27–3.33 in ♀), THL = 1.17 mm (1.00–1.33, 1.00–1.33), WL = 2.67 mm (2.27–2.83, 2.33–3.17), WW = 1.07 mm (0.90–1.13, 1.00–1.27).

Etymology. The name means “the Almighty” in the language of the Va ethnic group in Yunnan.

Stegana (Oxyphortica) setipes Wang & Chen, sp. nov.

urn:lsid:zoobank.org:act:57CE9B92-7CF3-4CD6-89A0-1E1C38CC0E9D

(Figs. 4D, 5N, 16)

Type specimen. Holotype, ♂ (SCAU, no. 111304), CHINA: Hesong, Menghai, Yunnan, 21°50′08″N, 100°05′51″E, 1,700–1,900 m a.s.l., 27.iii.2011, ex tussock (J. Lu).

Diagnosis. This species somewhat resembles S. (O.) tsauri sp. nov. in the male terminalia, but can be distinguished from the latter by the foretarsi first and second with two rows of ca. 30 long, recurvate fringe-like setae on anterior surface (Fig. 4D); abdominal tergites yellow medially, light brown laterally (Fig. 5N).

Description. Arista with five dorsal and three ventral branches. Clypeus brown. Mesoscutum and scutellum yellow. Katepisternum brownish above, yellow below. Male terminalia (Fig. 16): Epandrium with ca. 25 setae on posterior portion per side. Surstylus with two prensisetae dorsally, and numerous setae submedially and ventrally. Pregonites present. Aedeagus with several setae medially and numerous minute serrated processes laterally, fused with aedeagal sheaths dorsomedially. Aedeagal sheaths curved posteriorly, acute apically.

Measurements. BL = 2.33 mm in holotype, THL = 1.00 mm, WL = 2.03 mm, WW = 0.97 mm.

Etymology. A combination of the Latin words: seta + pes, referring to the foretarsi with long, fringe-like setae.

Stegana (Oxyphortica) shangrila Zhang & Chen, sp. nov.

urn:lsid:zoobank.org:act:3A1E56C9-47B2-481E-86D7-59827175DE19

(Figs. 4E, 5O, 17)

Type specimens. Holotype, ♂ (SCAU, no. 121372), CHINA: Hutiaoxia, Xianggelila, Yunnan, 27°11′24″N, 100°02′54″E, 1,700 m a.s.l., 22.viii.2011, ex tussock (H. Chen, J. Gao). Paratypes, CHINA: 2♂, 3♀ (1♂, 1♀ in KIZ, 0090655, 56; 1♂, 2♀ in SCAU, nos 121373–75), same data as holotype.

Diagnosis. Foretarsi first to third with a row of ca. 10 long, slightly valgus fringe-like setae on anterior surface (Fig. 4E). Aedeagus distally bifurcated, with numerous long and short setae (Figs. 17C, 17D); aedeagal sheaths distally triangular, strongly protruded posteriorly, plicated laterally, anterolaterally elongated, curved and fused with phallapodeme (Figs. 17C, 17D).

Description. Arista with seven to eight dorsal and three to five ventral branches. Clypeus brown. Mesoscutum, katepisternum and scutellum yellow. Abdominal first to fourth tergites yellow medially, brown to dark brown laterally; fifth and sixth dark brown, with yellow anteromedially (Fig. 5O). Sternites entirely yellow. Male terminalia (Fig. 17): Epandrium with ca. 19 setae on posterior portion per side. Cercus with pubescence dorsally. Surstylus with one stout prensiseta dorsally. Pregonites absent. Aedeagal sheaths elongated distolaterally, fused with phallapodeme.

Measurements. BL = 2.67 mm in holotype (2♂ and 3♀ paratypes: 2.53 in ♂, 3.00–3.13 in ♀), THL = 1.07 mm (1.07, 1.20–1.27), WL = 2.20 mm (2.13, 2.53–2.87), WW = 1.07 mm (1.07, 1.20–1.60).

Etymology. From the Tibetan lection, meaning an enchanting place.

Stegana (Oxyphortica) tsauri Zhang & Chen, sp. nov.

urn:lsid:zoobank.org:act:420D7B97-DDCC-4C46-B937-380DEECA2DF5

(Figs. 4F, 5P, 18)

Type specimens. Holotype, ♂ (SCAU, no. 122714), CHINA: Dabang, Chiayi, Taiwan, 23°28′47″N, 120°39′11″E, 630–800 m a.s.l., 15.x.2012, ex tree trunk (H. Chen). Paratypes, CHINA: 1♂, 1♀ (SCAU, nos 122715, 16), same data as holotype.

Diagnosis. This species differs from S. (O.) mengbalanaxi sp. nov. in having the foretarsi first and second with two to three rows of ca. 28 long, recurvate fringe-like setae, and first to fourth subapically each with one short, valgus fringe-like seta on anterior surface (Fig. 4F); abdominal first tergite brown; second medially brown, laterally black; third mostly black, brown along anterior margin; fourth to sixth black (Fig. 5P).

Description. Arista with five dorsal and three to four ventral branches. Clypeus black. Mesoscutum and scutellum brown. Katepisternum brownish. Abdominal sternites nearly brown. Male terminalia (Fig. 18): Epandrium with ca. 29 setae on posterior portion per side. Surstylus with one ventral and two dorsal prensisetae, and numerous setae submedially and ventrally. Pregonites absent. Aedeagus with several setae medially and numerous minute serrated processes laterally, fused with aedeagal sheaths dorsomedially. Aedeagal sheaths curved posteriorly, acute apically.

Measurements. BL = 2.73 mm in holotype (1♂ and 1♀ paratypes: 2.47 in ♂, 2.80 in ♀), THL = 1.13 mm (1.07, 1.13), WL = 2.67 mm (2.60, 3.00), WW = 1.07 mm (0.93, 1.13).

Etymology. The species is named in honor of Dr. Shun-Chern Tsaur (National Taiwan University) for helping HW Chen′s research in Taiwan.

Stegana (Oxyphortica) valleculata Zhang & Chen, sp. nov.

urn:lsid:zoobank.org:act:56D44D2D-0311-4A1A-B983-A0374959F113

(Figs. 4G, 5Q, 19)

Type specimens. Holotype, ♂ (SCAU, no. 123005), CHINA: Muyiji Park, Ximeng, Yunnan, 22°37′15″N, 99°35′42″E, 1,100–1,200 m a.s.l., 2.iv.2011, ex tussock (J. Lu). Paratypes, CHINA: 3♀ (SCAU, nos 123006–08), same data as holotype; 1♂, 1♀ (SCAU, nos 123009, 10), Yixiang, Pu′er, Yunnan, 22°44′19″N, 101°07′01″E, 1,420 m a.s.l., 12.xii.2012, ex tussock (J. Gao); 2♀ (SCAU, nos 111724, 25), Muyiji Park, Ximeng, Yunnan, 22°37′15″N, 99°35′42″E, 1,100–1,200 m a.s.l., 16.iv.2018, ex tree trunk, (L Gong); 1♂ (SCAU, no. 111437), Guanlei, Mengla, Yunnan, 22°06′26″N, 100°45′46″E, 650–870 m a.s.l., 24.iv.2016, ex tussock (J Huang).

Diagnosis. This species resembles S. (O.) shangrila sp. nov. in the aedeagus bifurcated, with short setae (Figs. 19C, 19D), but can be distinguished from the latter by the foretarsi first to third with two rows of ca. 27 long, slightly curved fringe-like setae on anterior surface (Fig. 4G), and surstylus with two stout prensisetae dorsally (Fig. 19B).

Description. Arista with five to six dorsal and three to four ventral branches. Clypeus brown. Mesoscutum and scutellum yellow. Katepisternum yellowish. Abdominal first tergite yellow; second and third tergites yellow medially, brown laterally; fourth and fifth yellow medially and brown along posterior margins and laterally in male, nearly brown to dark in female; sixth brown to dark brown (Fig. 5Q). Sternites yellow. Male terminalia (Fig. 19): Epandrium with ca. 19 setae on posterior portion per side. Surstylus with a row of numerous setae submedially and ca. seven setae ventrally. Pregonites absent. Aedeagal sheaths distolaterally elongated, curved and fused with phallapodeme.

Measurements. BL = 3.07 mm in holotype (1♂ and 4♀ paratypes: 2.97 in ♂, 2.73–3.17 in ♀), THL = 1.07 mm (1.10, 1.07–1.27), WL = 2.73 mm (2.90, 2.53–3.07), WW = 1.06 mm (1.10, 1.06–1.20).

Etymology. From the Latin word: valleculatus, referring to that the fly hides in the valley.

Stegana (Oxyphortica) wanhei Zhang & Chen, sp. nov.

urn:lsid:zoobank.org:act:D00C2F60-7938-461C-A234-B48C76A40FB2

(Figs. 4H, 5R, 20)

Type specimens. Holotype, ♂ (SCAU, no. 111202), CHINA: Hutiaoxia, Xianggelila, Yunnan, 27°11′24″N, 100°02′54″E, 1,700 m a.s.l., 22.viii.2011, ex tussock (H. Chen). Paratypes, CHINA: 48♂, 28♀ (10♂, 10♀ in KIZ, nos 0090625–44; 38♂, 18♀ in SCAU, nos 111203–58), ex tussocks and tree trunks, same data as holotype (H. Chen, J. Gao); 4♀ (SCAU, nos 121376–79), Menglun, Mengla, Yunnan, 21°59′57″N, 101°02′50″E, 820–890 m a.s.l., 24–26.xii.2003, ex tree trunks (H. Chen); 2♀ (SCAU, nos 121380, 81), Niuluohe, Jiangcheng, Yunnan, 22°30′N, 101°53′E, 1,000 m a.s.l., 21.iv.2010, ex tussock (L Wang); 1♂, 1♀ (SCAU, nos 122931, 32), Wuliangshan, Jingdong, Yunnan, 24°17′27″N, 100°19′48″E, 1,700–2,100 m a.s.l., 1.ix.2014, ex tussock (J. Gao); 2♂, 1♀ (SCAU, nos 122933–35), Xianheping, Anlong, Guizhou, 24°59′36″N, 105°36′14″E, 1,500 m a.s.l., 16.vii.2014, ex tussock (Y Zhang); 1♀ (SCAU, no. 122936), Wuliangshan, Jingdong, Yunnan, 24°17′27″N, 100°19′48″E, 1,700–2,100 m a.s.l., 16.vii.2009, ex tussock (L Wang); 1♂, 6♀ (SCAU, nos 111259–65), Qimaba, Lüchun, Yunnan, 22°49′03″N, 102°17″39″E, 1,280 m a.s.l., 30.x.2016, ex tussock (H. Chen); 4♂, 2♀ (SCAU, nos 111431–36), Nanling, Lancang, Yunnan, 22°47′57″N, 99°55′11″E, 1,800 m a.s.l., 12.viii.2016, ex tussock (H. Chen, L Gong).

Diagnosis. This species differs from S. (O.) gelea sp. nov. and S. (O.) jinmingi sp. nov. in having the apical processes of aedeagal sheaths curved in lateral view (Fig. 20D), abdominal fifth and sixth tergites yellow medially, narrowly black along lateral margin (Fig. 5R), and foretarsi first subapically with two rows of ca. 10 recurvate fringe-like setae, second with two short fringe-like setae on anterior surface (Fig. 4H).

Description. Arista with seven to nine dorsal and four to five ventral branches. Clypeus brown. Mesoscutum and scutellum brownish yellow. Katepisternum brown anteriorly, yellow posteriorly. Abdominal first tergite yellow; second to fourth tergites yellow medially, black laterally (Fig. 5R). Sternites yellow. Male terminalia (Fig. 20): Epandrium with ca. 35 setae on posterior portion per side. Surstylus with two prensisetae dorsally and two irregular rows of numerous setae submedially, acute ventrally. Aedeagus with several setae medially, mediodorsally fused with aedeagal sheaths, bifurcated distally. Aedeagal sheaths bifurcated distally.

Measurements. BL = 3.00 mm in holotype (5♂ and 5♀ paratypes: 2.87–3.40 in ♂, 3.53–4.47 in ♀), THL = 1.40 mm (1.40–1.60, 1.33–1.67), WL = 2.67 mm (2.87–3.00, 3.00–3.40), WW = 1.27 mm (1.27–1.47, 1.40–1.60).

Etymology. The species is named in honor of Mr. Wanhe Zhang (KIZ), a very capable driver in our multi-sited fieldworks.

Stegana (Oxyphortica) yangjin Zhang & Chen, sp. nov.

urn:lsid:zoobank.org:act:2316FD6E-5ED4-43EA-A3B3-1F48275BE2F8

(Figs. 4I, 5S, 21)

Type specimen. Holotype, ♂ (SCAU, no. 121371), CHINA: Beibeng, Motuo, Xizang, 29°14′36″N, 95°19′56″E, 1,100 m a.s.l., 2.x.2010, ex tussock (J. Gao).

Diagnosis. This species differs from S. (O.) mengbalanaxi sp. nov. in having the foretarsi first and second with two rows of ca. 12 long, slightly recurvate fringe-like setae on anterior surface (Fig. 4I), and abdominal tergites brown on first to third, dark brown on fourth to sixth (Fig. 5S).

Description. Arista with five dorsal and four ventral branches. Clypeus brownish yellow. Mesoscutum and scutellum yellow. Katepisternum yellow, slightly brown anteriorly. Abdominal sternites yellow. Male terminalia (Fig. 21): Epandrium with ca. 19 setae on posterior portion per side. Surstylus with two prensisetae dorsally, pubescence medially and setae submedially and ventrally. Pregonites absent. Aedeagus with several setae medially and numerous minute serrated processes laterally, fused with aedeagal sheaths dorsomedially. Aedeagal sheaths curved posteriorly, acute apically.

Measurements. BL = 2.84 mm in holotype, THL = 1.42 mm, WL = 2.34 mm, WW = 0.98 mm.

Etymology. A girl name from the Tibetan, meaning “her singing as though heavenly sounds in the full-moonlit starry sky”.

Stegana (Oxyphortica) hypophaia Zhang & Chen, sp. nov.

urn:lsid:zoobank.org:act:072AD3E7-F24B-4605-83EF-1CF9E9FA41D8

(Figs. 4J; 5T; 22)

Type specimens. Holotype, ♂ (SCAU, no. 123021), CHINA: Xinling, Badong, Hubei, 31°02′24″N, 110°20′48″, 650 m a.s.l., 7.x.2014, ex tussock (J. Gao). Paratypes, CHINA: 1♂, 3♀ (SCAU, nos 123022–24, 111266), same data as holotype.

Diagnosis. This species somewhat resembles S. (O.) crypta sp. nov. in some characters of the male terminalia (Figs 10, 22), but can be distinguished by the foretarsi first to fourth subapically each with one short, valgus fringe-like seta on anterior surface (Fig. 4J), aedeagal sheaths anterolaterally fused to phallapodeme, and aedeagus distally oblong and plicated (Figs. 22C, 22D).

Description. Arista with six to eight dorsal and four to five ventral branches. Clypeus black. Mesoscutum yellow anteriorly, brownish yellow posteriorly, with three thin, brownish, obscure stripes medially and sublaterally. Katepisternum yellow. Scutellum brown. Abdominal first tergite yellow; second to sixth tergites yellow medially, brown to dark brown laterally and posteriorly (Fig. 5T). Sternites yellow. Male terminalia (Fig. 22): Epandrium with ca. 20 setae on posterior portion per side. Cercus with sparse pubescence. Surstylus with two stout prensisetae dorsally, numerous setae medially and a few setae ventrally. Pregonites present.

Measurements. BL = 3.17 mm in holotype (1♂ and 2♀ paratypes: 2.58 in ♂, 2.27–3.13 in ♀), THL = 1.20 mm (1.17, 0.87–1.25), WL = 2.23 mm (2.06, 1.90–2.37), WW =1.13 mm (0.93, 0.97–1.00).

Etymology. A combination of the Greek words: hypos (=under) and phaios (gray), referring to the gray wing.